Faculty Publications

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Open Access
Carbon dynamics and estimates of primary production by harvest, 14C dilution, and 14C turnover
(Colorado State University. Libraries, 1992-04) Lauenroth, W. K., author; Milchunas, D. G., author; Ecological Society of America, publisher
Large plots of native shortgrass steppe were labeled with 14C to assess short-term patterns of carbon allocation and the long-term process of herbivory, death, and decomposition, and to compare estimates of net aboveground, crown, and root primary production using 14C dilution, 14C turnover, and traditional harvest methods. Stabilization of labile 14C via translocation, incorporation into structural tissue, and respiration and exudation required one growing season. Exudation was 17% of plant 14C after stabilization. Estimates of turnover time for leaves, crowns, and roots by 14C turnover were 3, 5, and 8 yr, respectively, yielding estimates of belowground production that were much lower than previously thought. Estimates of aboveground production by 14C turnover were close to those obtained by harvest of peak-standing crop, but lower than reported values obtained by harvest maxima-minima. Estimates of root production by harvest maxima-minima were zero in 2 of 4 yr. 14C turnover appeared to provide reliable estimates of aboveground, crown, and root production. In contrast to reliable estimates by 14C turnover, 14C dilution estimates of root production were anomalous. The anomalous estimates were attributed to a nonuniform labeling of tissue age classes resulting in differential decomposition/herbivory of 14C:12C through time, as well as movement and loss of labile 14C through the first growing season. Isotope-dilution methodologies may be unreliable for any estimate of pool turnover when the labeling period is not as long as pool-turnover time. Problems and biases associated with traditional harvest maxima-minima methods of estimating aboveground primary production are well known, but are greatly exacerbated when the method is used to estimate root production. Estimates of root production by 14C dilution were unrealistic. 14C turnover methodology provided reliable estimates of production in this community.
Open Access
CO2 enhances productivity, alters species composition, and reduces digestibility of Shortgrass Steppe vegetation
(Colorado State University. Libraries, 2004-01) Morgan, Jack A., author; Mosier, Arvin R., author; Milchunas, Daniel G., author; LeCain, Daniel R., author; Nelson, Jim A., author; Parton, William J., author; Ecological Society of America, publisher
The impact of increasing atmospheric CO2 concentrations has been studied in a number of field experiments, but little information exists on the response of semiarid rangelands to CO2, or on the consequences for forage quality. This study was initiated to study the CO2 response of the shortgrass steppe, an important semiarid grassland on the western edge of the North American Great Plains, used extensively for livestock grazing. The experiment was conducted for five years on native vegetation at the USDA-ARS Central Plains Experimental Range in northeastern Colorado, USA. Three perennial grasses dominate the study site, Bouteloua gracilis, a C4 grass, and two C3 grasses, Pascopyrum smithii and Stipa comata. The three species comprise 88% of the aboveground phytomass. To evaluate responses to rising atmospheric CO2, we utilized six open-top chambers, three with ambient air and three with air CO2 enriched to 720 mmol/mol, as well as three unchambered controls. We found that elevated CO2 enhanced production of the shortgrass steppe throughout the study, with 41% greater aboveground phytomass harvested annually in elevated compared to ambient plots. The CO2-induced production response was driven by a single species, S. comata, and was due in part to greater seedling recruitment. The result was species movement toward a composition more typical of the mixed-grass prairie. Growth under elevated CO2 reduced the digestibility of all three dominant grass species. Digestibility was also lowest in the only species to exhibit a CO2-induced production enhancement, S. comata. The results suggest that rising atmospheric CO2 may enhance production of lower quality forage and a species composition shift toward a greater C3 component.
Open Access
Hydrologic regime and herbivory stabilize an alternative state in Yellowstone National Park
(Colorado State University. Libraries, 2007) Wolf, Evan C., author; Cooper, David J., author; Hobbs, N. Thompson, author; Ecological Society of America, publisher
A decline in the stature and abundance of willows during the 20th century occurred throughout the northern range of Yellowstone National Park, where riparian woody-plant communities are key components in multiple-trophic-level interactions. The potential causes of willow decline include climate change, increased elk browsing coincident with the loss of an apex predator, the gray wolf, and an absence of habitat engineering by beavers. The goal of this study was to determine the spatial and temporal patterns of willow establishment through the 20th century and to identify causal processes. Sampled willows established from 1917 to 1999 and contained far fewer young individuals than was predicted from a modeled stable willow population, indicating reduced establishment during recent decades. Two hydrologically distinct willow establishment environments were identified: fine grained beaver pond sediments and coarse-grained alluvium. Willows established on beaver pond sediment earlier in time, higher on floodplain surfaces, and farther from the current stream channel than did willows on alluvial sediment. Significant linear declines from the 1940s to the 1990s in alluvial willow establishment elevation and lateral distance from the stream channel resulted in a much reduced area of alluvial willow establishment. Willow establishment was not well correlated with climate-driven hydrologic variables, but the trends were consistent with the effects of stream channel incision initiated in ca. 1950, 20-30 years after beaver dam abandonment. Radiocarbon dates and floodplain stratigraphy indicate that stream incision of the present magnitude may be unprecedented in the past two millennia. We propose that hydrologic changes, stemming from competitive exclusion of beaver by elk over browsing, caused the landscape to transition from a historical beaver-pond and willow mosaic state to its current alternative stable state where active beaver dams and many willow stands are absent. Because of hydrologic changes in streams, a rapid return to the historical state may not occur by reduction of elk browsing alone. Management intervention to restore the historical hydrologic regime may be necessary to recover willows and beavers across the landscape.
Open Access
Inertia in plant community structure: state changes after cessation of nutrient-enrichment stress
(Colorado State University. Libraries, 1995-05) Lauenroth W. K., author; Milchunas, D. G., author; Ecological Society of America, publisher
Water, nitrogen, and water-plus-nitrogen at levels beyond the range normally experienced by shortgrass steppe communities were applied from 1971 through 1975, plant populations were sampled through 1977, and the results of the experiment were published. Upon revisiting the plots in 1982, we found it apparent that large changes had occurred since 1977. Sampling was re-established in 1982 to follow trajectories of recovery. Our purposes in this paper are to examine how conclusions from this study changed through time, and discuss implications of these changes for monitoring potentially stressed ecosystems. Although productivities increased, dissimilarities in plant species composition at the end of the 5 year of nutrient treatments were not significantly different from controls. Two years after cessation of the treatments exotic "weed" species were increasing in water plus-nitrogen treated communities, and community dissimilarities were diverging in water and water-plus-nitrogen treated communities. Seven years after cessation of treatments all communities were significantly different from controls. Exotics were more than ten times more abundant in water-plus-nitrogen and nitrogen treated communities than they had been2 year post-treatment. A consistent trend in recovery of all treated communities was evident over the next 5 yr. However, the trend towards recovery reversed over the next four consecutive years in the previously water-plus-nitrogen and water treated communities. The four-to-five year cycles in species composition and abundance of exotics towards, and then away from, conditions in undisturbed control communities were not related to weather, but large accumulations of litter suggested biotic regulation. Inertia of existing plant populations, or the tendency to continue to occupy a site when conditions become unfavorable, can mask both future deterioration in ecosystem condition and unstable behavior resulting from environmental stressors. Time lags in initial response means that an ecosystem can pass a threshold leading to transitions to alternate states before it is evident in structural characteristics such as species composition. Global climate change and sulfur and nitrogen oxide pollutants also have the potential to act as enrichment-stressors with initial time lags and/or positive effects and cumulative, subsequent negative effects, rather than as disturbance forces with immediate negative impacts. Sociopolitical systems, however, often require change in biological variables or negative impacts before acting to ameliorate environmental problems. The manner in which conclusions changed at various periods in time, and the potential for time lags in responses of species populations, raises questions about which variables are most useful for detection of stress and how long studies must last to be useful.
Open Access
Interactions between individual plant species and soil nutrient status in Shortgrass Steppe
(Colorado State University. Libraries, 1995-06) Burke, Ingrid C., author; Vinton, Mary Ann, author; Ecological Society of America, publisher
The effect of plant community structure on nutrient cycling is fundamental to our understanding of ecosystem function. We examined the importance of plant species and plant cover (i.e., plant covered microsites vs. bare soil) on nutrient cycling in shortgrass steppe of northeastern Colorado. We tested the effects of both plant species and cover on soils in an area of undisturbed shortgrass steppe and an area that had undergone nitrogen and water additions from 1971 to 1974, resulting in significant shifts in plant species composition. Soils under plants had consistently higher C and N mineralization rates and, in some cases, higher total and microbial C and N levels than soils without plant cover. Four native grasses, one sedge, and one shrub differed from one another in the quantity and quality of above- and belowground biomass but differences among the six species in soil nutrient cycling under their canopies were slight. However, soils under bunchgrasses tended to have higher C mineralization and microbial biomass C than soil under the rhizomatous grass, Agropyron smithii. Also, the one introduced annual in the study, Kochia scoparia, had soils with less plant-induced heterogeneity and higher rates of C and N mineralization as well as higher levels of microbial biomass C than soils associated with the other species. This species was abundant only on plots that had received water and nitrogen for a 4-yr period that ended 20 year ago, where it has persisted in the absence of resource additions for 20 yr, suggesting a positive feedback between plant persistence and soil nutrient status. Plant cover patterns had larger effects on ecosystem scale estimates of soil properties than the attributes of a particular plant species. This result may be due to the semiarid nature of this grassland in which plant cover is discontinuous and decomposition and nutrient availability are primarily limited by water, not by plant species-mediated characteristics such as litter quality. That local plant-induced patterns in soil properties significantly affected ecosystem scale estimates of these properties indicates that consideration of structural attributes, particularly plant cover patterns, is critical to estimates of ecosystem function in shortgrass steppe.
Open Access
Relationships between groundwater use, water table, and recovery of willow on Yellowstone's northern range
(Colorado State University. Libraries, 2011) Johnston, Danielle B., author; Cooper, David J., author; Hobbs, N. Thompson, author; Ecological Society of America, publisher
Excessive levels of herbivory, incision of stream channels, and climate warming are believed to be responsible for the decline of woody deciduous plants in riparian zones in western North America, declines that are likely to be associated with diminished biological diversity. In the northern elk wintering range of Yellowstone National Park, USA, over browsing by elk (Cervus elaphus), lowered water tables resulting from stream incision, and loss of activity by beaver (Castor canadensis) have been implicated in the decline of willow (Salix sp.) communities. Reducing elk browsing appears sufficient for willow recovery in some areas, but where water table changes have been dramatic, recovery may be slow or absent. The importance of water table changes is disputed because experimental results demonstrate water table limitations, but water table depth has failed to explain variation in willow height at landscape scales. One explanation for this apparent discrepancy is that willows that have survived intensive browsing by elk have maintained access to groundwater despite declining water tables. Using stable isotopes of water, we examined the relationships between groundwater use, water table depth, and height of heavily browsed Salix geyeriana. Salix geyeriana groundwater use varied from 30% to 80%, and was higher later in the growing season, when soil water was less available and shoot water potentials were lower. Late season groundwater use explained 26% of the variation in total height of willows (P = 0.002), with taller plants using more groundwater. Water table depth explained only 8% of the variability in total height (P=0.051), with shorter willows having deeper water table depths. Groundwater use and water table depth were uncorrelated. Height recovery following a winter of heavy browsing was related to groundwater use, but not groundwater depth. We suggest that access to deeper water sources alleviates late season water stress, allowing for more rapid height recovery and higher total plant height. Variability in groundwater access may account for variability in height recovery at landscape scales.
Open Access
Soil organic matter recovery in semiarid grasslands: implications for the Conservation Reserve Program
(Colorado State University. Libraries, 1995-08) Coffin, Debra P., author; Lauenroth, William K., author; Burke, Ingrid C., author; Ecological Society of America, publisher
Although the effects of cultivation on soil organic matter and nutrient supply capacity are well understood, relatively little work has been done on the long-term recovery of soils from cultivation. We sampled soils from 12 locations within the Pawnee National Grasslands of northeastern Colorado, each having native fields and fields that were historically cultivated but abandoned 50 years ago. We also sampled fields that had been cultivated for at least 50 years at 5 of these locations. Our results demonstrated that soil organic matter, silt content, microbial biomass, potentially mineralizable N, and potentially respirable C were significantly lower on cultivated fields than on native fields. Both cultivated and abandoned fields also had significantly lower soil organic matter and silt contents than native fields. Abandoned fields, however, were not significantly different from native fields with respect to microbial biomass, potentially mineralizable N, or respirable C. In addition, we found that the characteristic small-scale heterogeneity of the shortgrass steppe associated with individuals of the dominant plant, Bouteloua gracilis, had recovered on abandoned fields. Soil beneath plant canopies had an average of 200 g/m2 more C than between-plant locations. We suggest that 50 years is an adequate time for recovery of active soil organic matter and nutrient availability, but recovery of total soil organic matter pools is a much slower process. Plant population dynamics may play an important role in the recovery of shortgrass steppe ecosystems from disturbance, such that establishment of perennial grasses determines the rate of organic matter recovery.
Open Access
Water tables constrain height recovery of willow on Yellowstone's northern range
(Colorado State University. Libraries, 2008) Bilyeu, Danielle M., author; Cooper, David J., author; Hobbs, N. Thompson, author; Ecological Society of America, publisher
Excessive levels of herbivory may disturb ecosystems in ways that persist even when herbivory is moderated. These persistent changes may complicate efforts to restore ecosystems affected by herbivores. Willow (Salix spp.) communities within the northern range in Yellowstone National Park have been eliminated or degraded in many riparian areas by excessive elk (Cervus elaphus L.) browsing. Elk browsing of riparian willows appears to have diminished following the reintroduction of wolves (Canis lupis L.), but it remains uncertain whether reduced herbivory will restore willow communities. The direct effects of elk browsing on willows have been accompanied by indirect effects from the loss of beaver (Castorcanadensis Kuhl) activity, including incision of stream channels, erosion of fine sediments, and lower water tables near streams historically dammed by beaver. In areas where these changes have occurred, lowered water tables may suppress willow height even in the absence of elk browsing. We conducted a factorial field experiment to understand willow responses to browsing and to height of water tables. After four years of protection from elk browsing, willows with ambient water tables averaged only 106 cm in height, with negligible height gain in two of three study species during the last year of the experiment. Willows that were protected from browsing and had artificially elevated water tables averaged 147 cm in height and gained 19 cm in the last year of the experiment. In browsed plots, elevated water tables doubled height gain during a period of slightly reduced browsing pressure. We conclude that water availability mediates the rate of willow height gain and may determine whether willows grow tall enough to escape the browse zone of elk and gain resistance to future elk browsing. Consequently, in areas where long-term beaver absence has resulted in incised stream channels and low water tables, a reduction in elk browsing alone may not be sufficient for recovery of tall willow stands. Because tall willow stems are important elements of habitat for beaver, mitigating water table decline may be necessary in these areas to promote recovery of historical willow–beaver mutualisms.

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